Variation in floristic and trait composition along environmental gradients in the herb layer of temperate forests in the transition zone between Central and SE Europe

Species- and trait-environment linkages in forest plant communities continue to be a frequent topic in ecological research. We studied the dependence of floristic and functional trait composition on environmental factors, namely local soil properties, overstory characteristics, climatic parameters and other abiotic and biotic variables. The study area comprised 50 monitoring plots across Slovenia, belonging to the EU ICP Forests monitoring network. Vegetation was surveyed in accordance with harmonized protocols, and environmental variables were either measured or estimated during vegetation sampling. Significant predictors of species composition were identified by canonical correspondence analysis. Correlations between plant traits, i.e. plant growth habit, life form, flowering features and CSR signature, were examined with fourth-corner analysis and linear regressions. Our results show that variation in floristic composition was mainly explained by climatic parameters (mean annual temperature, mean annual precipitation), soil properties (pH) and tree layer-dependent light conditions. Trait composition was most closely related with tree layer characteristics, such as shade-casting ability (SCA, a proxy for light availability in the understory layer), tree species richness and tree species composition. Amongst soil properties, total nitrogen content and soil texture (proportion of clay) were most frequently correlated with different species traits or trait states. The CSR signature of herb communities was associated with tree layer SCA, soil pH and mean annual temperature. The floristic composition of the studied herb-layer vegetation depended on temperature and precipitation, which are likely to be influenced by ongoing climate change (warming and drying). Trait composition exhibited significant links to tree layer characteristics and soil conditions, which are in turn directly modified by forest management interventions